BASTERIA, 63: 139-163, 1999
Notes on the non-marine molluscs of the island of Borneo 9. The
and genera Cyclophorus, Leptopoma , Craspedotropis (Gastropoda Prosobranchia: Cyclophoridae)
J.J. Vermeulen
Nationaal Natuurhistorisch Museum, P.O. Box 9517, NL 2300 RA Leiden, The Netherlands
The species of the generaCyclophorus,Leptopoma, and Craspedotropis (Cyclophoridae) occurring
is from the on Borneo are revised. The genus Craspedotropis recorded Borneo for first time,
including two new species (C. juvenilis and C. andrei), and one species previously included in
Jerdonia (C. borneensis).
Key words: Gastropoda, Prosobranchia, Cyclophoridae, Cyclophorus, Leptopoma, Craspedotropis,
taxonomy, Malaysia, Indonesia, Borneo.
INTRODUCTION
The revision of the largest family of Bornean land snails, the Diplommatinidae, being completed in the previous instalment of the series (Vermeulen, 1996), the author has started a revision of the second largest family occurring on Borneo, the Cyclophoridae. of The first results are presented in this paper. It provides a revision the cyclophorid 1810 1847 and genera Cyclophorus Montfort, (4 taxa), Leptopoma Pfeiffer, (6 taxa), Craspe-
dotropis Blanford, 1864 (3 taxa). The first two include species with large or conspicuously coloured shells that the first be collected almost were among to on Borneo, now two
centuries ago. They were also among the first to be described. Unfortunately, the
descriptions and illustrations of these early names are often barely accurate enough to recognize the species, and, moreover, in some cases the type material has to be pre- accessible the author. Yet it is that the sumed lost, or was not to expected synonyms listed below species such as Cyclophorus perdix borneensis (Metcalfe, 1851),Leptopoma undatum L. and L. sericatum (Metcalfe, 1851), pellucidum (Grateloup, 1840), (Pfeiffer, 1851) are largely accurate, and that at least no other valid species remain hidden under the synonymized names
Access to the literature of this family is greatly facilitated by the various revisions and enumerations that have been published, the last of these being Kobelt (1902a, 1902b) and Gude (1921). Although these works are rather uncritical in many cases, they at least list ofthe nominal Borneo and in surround- provide a fairly complete taxa recognized on ing areas.
References to material in the collection of the author are abbreviated as: V, followed by a collection number. Most of this material, including all holotypes of species described here will be deposited in the Nationaal Natuurhistorisch Museum (RMNH). If available, will be distributed other institutes. For other collections the abbre- paratypes to following British collection viations are used: BMNH, Museum (Natural History), London; K, private
1 For no. 8 in this series see Basteria 60: 87-138, 1996. 140 BASTERIA, Vol. 63, No. 4-6, 1999 of Mr. K. Kittel (Germany); R, private collection ofDr. J. G. M. Raven (The Netherlands);
UF, The Florida Museum ofNatural History, Gainesville. After each reference to material seen by the author the number of specimens is given after a slash: /. few other abbreviations used in the references A are geographical in the text, mainly derived from the Indonesian language: Bt. = bukit (hill); G. = Gunung (mountain); K.
= Kalimantan of the Indonesian of abbreviation used in (the name part Borneo, only the names of the various provinces); Kpg. = Kampong (village); P. = Pulau (island); Sg.
= Sungei (River).
The drawings were made by the author, with pencil, using a Wild M 8 stereo micro- with lucida device. scope camera
SYSTEMATIC PART
Cyclophorus Montfort, 1810
Shell with whitish brownish large (30-50 mm wide), solid, opaque, a silky lustre, to pale or greenish yellow, usually marked with darker brown. Protoconch with well-spaced, thin radial riblets, and a fine spiral striation; transition towards the teleoconch clearly visible. Spire low conical or only slightly raised. Radial sculpture: growth lines mainly, sometimes with riblets close the Umbilicus inconspicuous, spaced to protoconch. open. Aperture circular to elliptic. Peristome thickened and reflected, angular edge without a sinus; peristome either simple, often consisting of thin, overlapping layers towards the
or double. Periostracum without hairs other outer margin, thin, smooth, or appendages. with Operculum thin, entirely corneous, many whorls, margins of the whorls not or hardly free, not raised. forest floor Ecology. Probably living on the (based on observations on extralimital species).
Distribution. India to Japan, southwards to Papua New Guinea, Indonesia. - Notes. Large genus, including about 200 species. Particularly well-represented from India to Indochina, and in The Philippines, relativelv few species on the Sunda shelf.
Key to the Bornean species of Cyclophorus
series of (check as large specimens as possible)
la The last 3/4 whorl distinctly but obtusely shouldered towards the suture. Peristome double in fully adult
the inner 7 from the 1- shells, protruding up to mm outer Cyclophorus niahensis
lb Either last whorl somewhat flattened towards the the last 3/4 whorl the 3/4 well-rounded,at most suture; or
shouldered towards the suture, and peristome simple 2
2a Spire depressed, apex only slightly raised. Suture channelled from the start ofthe teleoconch onwards. Spiral
2 - sculpture absent or nearly so Cyclophorus phlegethon
low conical. Suture but in the last whorl. 2b Spire depressed, not channelled, or only slightly so 1 1/2 Spiral
old 3 sculpture present, fine, wavy, but easily wearing off in specimens
towards the 3a Peristome either simple, consisting of thin, overlapping layers outer margin, or double, with
the inner to 2 mm from the outer. Umbilicus 7.5-9.0 mm wide peristome protruding up
3 - Cyclophorus kinabaluensis
3b Peristome simple, not consisting ofthin layers towards the outermargin (except sometimes in the umbilical
region) in fresh shells. Umbilicus 3.8-7.1 mm wide 4 - Cyclophorus borneensis Vermeulen: Non-marine molluscs Borneo & of (Cyclophorus, Leptopoma Craspedotropis) 141
1- Cyclophorus niahensis Godwin Austen, 1889 (fig. 1)
Cyclophorus niahensis Godwin Austen, 1889: 334; holotype ("Niah Hills") leg. Everett, BMNH 89.12.7.3.
Material SARAWAK. Div.: seen. 4th Kakus Mts., S. of Bintulu (leg. Bryant, UF
268445/1, see note below); G. Subis (Batu Niah) (leg. Dorman et al., UF 196446/3, do., UF 196448/3; leg. Everett, BMNH/3, see also above); G. Subis, W. of Batu Niah (leg. Raven, R/6); Rumah Malang (leg. Lopez, R 1607/1); Materae (leg. Lim Chan
Koon, V 5795/1). "North Borneo" (BMNH 1825/1). "Borneo" (RMNH/1).
Shell cream-coloured to pale brownish, with an irregular zig-zag pattern of darker brown, with a pale band at the periphery, with a wide band of dark (reddish) brown below and often above the periphery, partially dissolved into thin, more or less contin- lines umbilical whitish brown. low conical. uous spiral or not; region to pale Spire Whorls 4 1/4-4 7/8, the last 3/4 whorl distinctly, obtusely shouldered close to the with obtuse with suture, basally usually a distinct, spiral ridge, periphery a slight to distinct, obtuse edge. Suture depressed, somewhat channelled in the last 1 1/2 whorl. Umbilicus 6.3-8.5 covered the Spiral sculpture present, fine, wavy. mm wide, not by peristome. Peristome white, thickened and reflected, double, with the inner peristome 7 from the 22-30 width 37-50 protruding up to mm outer. Height mm, mm; height
16.5-23 width 18.5-28 mm. aperture mm, -- Ecology. Found in lowland forest on soils over limestone and sandstone/shale bedrock.
Distribution. Borneo: Sarawak, 4th Div., Upper Tatau Valley, Batu Niah and surroundings, Middle Baram Valley. shell from Tatau Notes. The only the Valley (UF 268445), a sub-adult with a thin peristome, lacks the spiral ridge on the basal surface.
2- Cyclophorus phlegethon Godwin Austen, 1889 (fig. 2)
Cyclophorus phlegethon Godwin Austen, 1889: 335; holotype ("Molu Hills") leg. Hose, BMNH 1998011.
everetti E. A. 1893: BMNH Cyclophorus Smith, 343; syntypes ("Barit Mountain") leg. Everett, (not seen).
Material seen. SARAWAK. 4th Div.: Bt. Pelamau (leg. Haegens, V 5613/1); G. Mulu G. Mulu National Tutoh area (leg. Everett, BMNH/1, see above); Park, valley, G. Benarat (leg. Wilford, UF 268448/2).
Shell but with of darkbrown whitish, an irregular zig-zag pattern usually leaving only of the with of the with wide patches ground colour, a row pale spots at periphery, a band ofdarkerbrown below the periphery; umbilical region whitish to pale brown. Spire raised. with depressed, apex only slightly Whorls 3 1/2-4, well-rounded, periphery or without a slight, obtuse edge. Suture distinctly channelled from the start of the teleo- conch onwards. absent Umbilicus 6.5-8.5 Spiral sculpture or nearly so. mm wide, not
covered by the peristome. Peristome dull whitish or pale brownish, only slightly thick- ened and reflected except on the columellar side; either simple, or double with the inner 2 from the 18.5-21 width 34-41 peristome protruding up to mm outer. Height mm, mm;
height aperture 15-18 mm, width 17.5-21 mm. Found in forest limestone Recorded from lowland Ecology. on or near outcrops. to up to 1200 m asl.
Distribution. Borneo: Sarawak, 4th Div., G. Mulu area and surroundings. 142 BASTERIA, Vol. 63, JVo. 4-6, 1999
Fig. 1. Cyclophorus niahensis Godwin Austen, 1889; Sarawak, G. Subis (RMNH) (shell height 22 mm). Vermeulen: Non-marine molluscs of Borneo (Cyclop/torus, Leptopoma & Craspedotropis) 143
C. Godwin Bt. Pelamau 22 Figs. 2-3. Cyclophorus spec. 2, phlegethon Austen, 1889; Sarawak, (V) (shell height mm). 3, C. kinabaluensis E. A. Smith, 1895; Sabah, Bt. Gomantong (V) (shell height 30 mm). 144 BASTERIA, Vol. 63, Mo. 4-6, 1999
3- Cyclophorus kinabaluensis E. A. Smith, 1895 (fig. 3)
Cyclophorus kina-baluensis E. A. Smith, 1895: 1 18; type ("Kina Balu, N. Borneo" BMNH (not seen).
Material seen. SABAH. Sandakan Zone: Bt. Gomantong, 30 km S. of Sandakan
(leg. Lee et al., UF 114724/6; V 1593/4). "Sabah" (leg. Beal-Maltbie, UF 242072/2). Dutch Borneo 1893-94, KALIMANTAN. Kalimantan Barat: G. Tilung (leg. exp. RMNH/1).
Shell white with an of to pale yellowish or greenish brown, irregular zig-zag pattern without less brown, often with a pale band at the periphery, with or thin, more or continuous spiral lines of darker brown above the periphery; below the periphery often thin umbilical with a band of darker brown partially dissolved similar, spiral lines; region whitish to pale brown. Spire low conical. Whorls 4 1/4-4 3/4, well-rounded, somewhat towards the with obtuse flattened suture, periphery a slight to distinct, to sharp edge.
Suture but not channelled. depressed, or hardly Spiral sculpture present, fine, wavy, easily wearing off in old specimens. Umbilicus 7.5-9.0 mm wide, not covered by the peristome. Peristome white to reddish, thickened and reflected; either simple, consisting ofthin, overlapping layers towards the outer margin, or double, with the inner peristome
2 mm from the outer. 25-28 width 43-47 protruding up to Height mm, mm; height aperture 18-21 mm, width 21-29 mm.
Ecology. In forest on various soils; recorded are limestone and granodiorite bed- rock. Recorded at sea level, probably at higher altitude in the Kinabalu area.
Distribution. Borneo: Sabah, G. Kinabalu, N. and E. coast. Kalimantan, K. Barat,
Kapuas basin.
4- Cyclophorus perdix borneensis (Metcalfe, 1851) (fig. 4)
Cyclostoma borneensis Metcalfe, 1851: 71; holotype ("Borneo") (not seen).
Cyclophorus borneensis; Pfeiffer, 1852: 63.
Cyclophorus cochranei Godwin Austen, 1889: 334; holotype ("Niah Hills") leg. Everett, BMNH 1889.12.7.5.
Cyclophorus cochranei var. ochraceus Godwin Austen, 1889: 335; holotype ("Busan Hills") leg. Everett, BMNH
1889.12.7.6.
Cyclophorus talboti Godwin Austen, 1889: 335; holotype ("Busan Hills") leg. Everett, BMNH 1889.12.7.7.
Cyclophorus perdix borneensis; Van Benthem Jutting, 1959: 69. See also Rensch, 1934: 739.
Material seen. SARAWAK. Ist Div.: Busau (leg. Beal-Maltbie, UF 242005/2; leg. of C. Everett, BMNH/1, see above, type cochranei var. ochraceus; do., BMNH/1, see above, of C. km 31.6 road UF G. type talboti); quarry Kuching-Bau (leg. Auffenberg, 183949/3);
Pangga 3 km E.N.E. of Bau (V 2072/3); near Bau (leg. De Vogel, V 2640/2 ); 3 km S.W.
of Taiton goldmine, near Bau (leg. Auffenberg, UF 183891/5); hill S. of Bau goldmine, Bt. Bau 2.1 km S. of Bau (leg. Auffenberg, UF 183836/4); Jagoi near (leg. Schuiteman, V 5474/1); G. Kapur 6 km S.E. of Bau (V 2044/3; do., 2045/7); Lobang Angin 2 km
S.W. ofBau (V 2081/4); Fairy Cave approx. 8 km S.W. ofBau (leg. Dorman, UF 196306/
2); G. Jambusan 4 km S.E. of Bau (V 2064/>lO ); G. Muan near kpg. Peninjau Lama,
22.5 km S.W. of Kuching (leg. Auffenberg, UF 183824/1; do., UF 183929/2); Hume Rock, 22.9 km S.E. ofKuching (leg. Auffenberg, UF 183998/1); Kpg. Beratok along road Kuching-Serian (leg. Auffenberg, UF 184003/5; do., UF 184026/10; V 2000/4); Semen-
goh Botanic Garden (leg. Rhea Warren, UF 28423/2; leg. De Vogel, V 2318/1); Kpg. Vermeulen: Non-marine molluscs of Borneo (Cyclophorus, Leptopoma & Craspedotropis) 145
Fig. 4a-d. Cyclophorus perdix borneensis (Metcalfe, 1851). a-b, Sarawak, G. Subis (V) (shell height 27 mm); c-d,
Sarawak, G. Kapur 6 km SE. of Bau (V) (shell height 27 mm). 146 BASTERL4, Vol. 63, No. 4-6, 1999
Tiang Bekap 10 km SS.W. of Kpg. Beratok (leg. De Vogel, V 2593/1); G. Wah S. of Braang, 23 miles S. of Kuching (leg. Wilford, UF 268446/4); G. Gayu 23 miles S. of
Kuching (leg. Wilford, UF 268450/3); Kpg. Segur Benuk, mile 21 Penrissen Road (leg. De Vogel, V 2558/5); G. Lelat 1 mile S.W. of Nyabet, 24 miles S.S.E. of Kuching (leg.
Wilford, UF 268449/1); 7.5 km W.N.W. of Kpg. Piching, near Serian (V 1900/8); G.
Selabor, W. of Kpg. Lobang Batu, 12.5 km S. of Tebakang (leg. Wilford, UF 268444/
5; V 2034/>10). 4th Div.: G. Subis (Batu Niah) (leg. Dorman, UF 196442/3; do., UF
196443/2; do., UF 196445/5; leg. Everett, BMNH/1, see above; leg. Holzmark, UF 24830/4; do, UF 24831/5; do, UF 24832/4; do, UF 24833/5; do, UF 24834/4; do,
UF 24835/5; leg. Sutton, UF 196405/1; leg. Wall, UF 268447/4; V 1580/>10); G. Subis,
W. ofBatu Niah (leg. Raven, R 1618/>10); G. Subis, 3 km W. of Batu Niah (leg. Raven, R/4); G. Subis, 4 km S. ofBatu Niah (leg. Raven, R L 1674/10); Laogan Bunut, S. edge
(leg. Raven, R L 1649/1). SABAH. West Coast Zone: G. Kinabalu (leg. Everett, UF KALIMANTAN. Kalimantan Barat: Dutch Borneo 161689/3). Liang Kubung (leg. exp. 1893-94, RMNH/1); Dadap Ketungau (do, RMNH/1); G. Sekedau (do, RMNH/1); G. Kenepai (do, RMNH/3); Sintang (do, RMNH/2); Sg. Sibau (do, RMNH/5); Pulau, Sg.
Sibau (do, RMNH/1). Kalimantan Selatan: G. Siamang near Desah Liu, 30 km E. of Muara limestone W. of the Tandjung (V 3088/7); Jaro near Uja, escarpment village (V
314-1 /> 10); limestone escarpments along road Benualawas-Limbungan (V 2905/1). Kalimantan Timur: Pt. ITCI logging concession (leg. Van Valkenburg & Galzin, V 4526/
1). "Borneo" (RMNH/1).
Shell white to pale yellowish brown, usually (albino specimens occur) with an irregular of dark often with of white below the and zig-zag pattern brown, a row spots suture at the periphery, without any thin, continuous spiral lines, but with one wide band or two narrower bands of darker brown below the periphery; umbilical region white, usually with fewer brown markings than elsewhere. Spire low conical. Whorls 4 1/2- sometimes somewhat flattened shouldered towards the 5, well-rounded, or suture, with without obtuse with keel. periphery or an to acute edge, or a Suture depressed, but in the last 1 1/2 whorl. not channelled, or only slightly so Spiral sculpture present, Umbilicus fine, wavy. 3.8-7.1 mm wide, sometimes partly covered by the peristome.
Peristome white or cream-coloured, slightly to distinctly thickened and distinctly reflect- ed, simple, not consisting of thin layers towards the outer margin (except sometimes in the umbilical in fresh shells. 19-29 width 30-46 region) Height mm, mm; height aperture
14-21 mm, width 16-28 mm.
Ecology. Primary forest on various soils, most numerous on limestone bedrock.
Also in secondary vegetation. Distribution. Borneo: widespread but in Sabah known from G. Kinabalu only. Also in W. Malaysia, and probably Singapore (see below). Notes. Next the above includes: C. to subspecies, Cyclophorus perdix p. perdix (Broderip
& Sowerby, 1830) Java to Bali), C. p. tuba (Sowerby, 1842) (Sumatra, W. Malaysia), C. p. aquila (Sowerby, 1843) (Singapore, W. Malaysia and possibly further North). Extral- intermediates between the latter and C. imitally, two p. borneensis occur, rendering the range of the Bornean subspecies uncertain. In of considerable spite a variability in shell colour, the number of riblets on the protoconch, presence/absence of a peripheral keel, and the width of the umbilicus, no further taxa can be distinguished within C. p. borneensis. Locally, states of these characters predominate, for instance at Batu Niah where most specimens display a comparatively umbilicus. narrow The surroundings of Batu Niah are inhabited by a form with a unusually sharp peripheral keel. Vermeulen: Non-marine molluscs of Borneo (Cyclophorus, Leptopoma & Craspedotropis) 147
Leptopoma Pfeiffer, 1847
Shell medium sized with whitish (9.5-24 mm wide), thin, a silky lustre, to corneous, with or without colour markings. Protoconch smooth or with fine spiral ridges, transition towards the teleoconch unmarked. Spire conical. Radial sculpture: growth lines mainly. often combined with threadlike Umbilicus Spiral sculpture: striation, ridges. open, or partly or entirely covered by the peristome. Aperture about circular. Peristome simple, more or less thickened and reflected, angular edge without a sinus; Periostracum thin, smooth, without hairs or other appendages. with of the whorls neither Operculum thin, entirely corneous, many whorls, margins free nor raised.
Distribution. India to Taiwan, southwards to Australia, eastwards to the Pacific. - Notes. Large genus, including over 100 species. Speciose in The Philippines, relatively few species on the Sunda shelf.
Specific delimitation is problematical in Leptopoma due to a general lack ofreliable shell characters, and to morphologically intermediate forms between several nominal species. A marked in shell within has caused further variability colouring many species compli- cations, because species names have been proposed mainly based on states of this character. Jonges (1980) studied the genitalia of various species of Leptopoma and found some differences; perhaps further anatomical research may lead to a better resolution of the into genus species. The is used below describe thickened often term 'lip', to a rim, present on the inner edge of the reflected portion of the peristome.
Key to the Bornean species of Leptopoma
(check as large series of specimens as possible)
la The first few whorls moderately convex, the others almost flat to slightly convex 2 lb All whorls moderately to distinctly convex 4
2a Palatal side of the peristome with a distinct lip that gradually increases in height towards the angular edge
in fully adult specimens 1 - Leptopoma trochus
2b Palatal side of the peristome without a lip; or peristome with a lip that gradually decreases in height towards
the angular edge 3
in 3a Periphery with a distinct keel that rather abruptly changes into a less conspicuous edge the last 1/3-1/4
of the keel when shell is whorl, thus disrupting the curvature the observed from below
2 - Leptopoma undatum
3b Periphery with a moderately distinct keel that may gradually change into a less conspicuous edge towards
the but without the curvature of the keel when the shell is observed from below aperture, disrupting ...
3 - Leptopoma bourguignati
4a (1) Reflected portion of the palatal peristome distinctly convex 4 - Leptopoma signatum
4b Reflected portion of the palatal peristome flat or only slightly convex (lip, if present not taken into
consideration) 5
5a Periphery rounded (rarely angular), at most with a thread-like, inconspicuous keel. Spiral sculpture: above
the often with five thread-like similar the keel but still periphery up to inconspicuous, ridges to peripheral
1 5 - weaker, below the periphery sometimes such a ridge Leptopoma pellucidum
5b Periphery rounded to angular, usually with a moderatelydistinct ridge. Spiral sculpture: above the periphery
3-6 rather distinct, thread-like ridges, below the periphery l-2(-6) such ridges 6 - Leptopoma sericatum 148 BASTERIA, Vol. 63, No. 4-6, 1999
1- Leptopoma trochus Dohrn, 1862 (fig. 5)
Leptopoma trochus Dohrn, 1862: 182; type ("in Maligi insulae Mindanao") leg. Semper (not seen)
Leptopoma geotrochiforme E. A. Smith, 1895: 119; syntypes ("Mount Rabong, Sarawak") BMNH 94.4.21.128-
9/2 & 94.7.20.85-6/2.
Material SARAWAK. seen. Ist Div.: G. Rabong (BMNH/4. see above; BMNH 1825/1; RMNH/1); Kpg. Tiang Bekap 10 km S.S.W. of Kpg. Beratok (leg. De Vogel, V 2595/2).
Shell white, with or without translucent banding parallel to the growth lines. Whorls 5 1 /4-5 the first few the others almost flat but 5/8, moderately convex, to slightly convex, the last also slightly to moderately concave towards the periphery; periphery with a distinct keel that continues the Suture between the first up to peristome. moderately depressed whorls, elsewhere slightly depressed. Spiral sculpture: a fine striation, often combinedwith 3-5 Umbilicus covered slightly raised, inconspicuous ridgelets. narrow, largely or entirely the Peristome the side by peristome. white, on palatal usually slightly thickened and with distinct that increases in towards the abruptly reflected, a lip gradually height angular and continues edge then on the parietal side as a somewhat lower lip again. Height 15- 19.5 width 17-22 5.5-9.5 width 9-12 mm, mm; height aperture mm, mm.
Ecology. Lowland forest on limestone soil. Distribution. Borneo: Sarawak, Ith Div. Also in The Philippines.
2- Leptopoma undatum (Metcalfe, 1851) (fig. 6)
Cyclostoma undatum Metcalfe, 1851: 71; type ("Borneo") (not seen)
Leptopoma undatum; Pfeiffer, 1852: 113.
Helix niahensis Godwin BMNH (Geotrochus) Austen, 1891: 44; type ("Niah Hills") leg. Everett, (not seen)
Leptopoma niahense; E. A. Smith, 1895: 119.
E. 1895: British Leptopoma skertchlyi A. Smith, 119; syntypes ("Mount Ambun, North Borneo") leg. Skcrtchly,
BMNH 95.1.16.1-2/2.
Papuina niahensis; Van Benthem Jutting, 1933: 58
Material -- SARAWAK. Ist Div.: km 31.6 road seen. quarry Kuching-Bau (leg. Auffenberg, UF 183947/2; do., UF 183986/1); G. Pangga 3 km E.N.E. of Bau (V 2150/ 2); 2-3 km SW. of Bau (leg. Dorman, UF 196276/1); 1 km S. of Bau, S. edge of active mine gold (leg. Dorman, UF 196368/1); Bt.Jagoi near Bau (leg. Schuiteman, V 5473/
3); Lobang Angin 2 km S.W. of Bau (V 2086/5); G. Muan near kpg. Peninjau Lama,
22.5 km S.W. of Kuching (leg. Auffenberg, UF 183825/2); Semengoh Botanic Garden (leg. De Vogel, V 2319/1). 4th Div.: G. Subis (Batu Niah) (leg. Dorman, UF 196452/
1; V 1544/1); G. Subis, limestone quarry (leg. Raven, R/3); Lambir oil palm plantation
(leg. Raven, R L 1603/1); Lambir Hills (leg. Sleumer, RMNH/1); do., near canteen (leg.
Raven, R 1586/1); do., near summit (leg. Raven, R 1588/1). BRUNEI. Temburong,
Labi Road 13 km from S. Liang (leg. De Vogel, V2494). SABAH. Interior Zone: 5.5 km N.N.E. of Simatuoh, 9 km E. of Sepulot (leg. Dorman, UF 196516/1). West Coast Zone: G. Kinabalu National Park (RMNH/2); do., Headquarters area (V 1195/2); do., Silau-Silau trail V trail towards (leg. Chan, 3647/1; do., top (V 1440/1); do., Poring
Hot Springs (leg. Kittel, K/l). Sandakan Zone: Tawai Mountains near Telupid (V Vermeulen: Non-marine molluscs of Borneo (Cyclophorus, & Craspedotropis) 149
4-7. and C. borneensis Beratok Figs. Cyclophoruss spec, Leptopoma spec. 4e, perdix (Metcalfe, 1851); Sarawak, Kpg.
(V) (shell height 32 mm). 5a-c, L. trochus Dohrn, 1862; Sarawak, Kpg. Tiang Bekap (V) (shell height 15 mm).
Sabah L. 6c, L. undatum (Metcalfe, 1851), (RMNH) (shell height 15 mm). 7c, bourguignati Issel, 1874, syntype
of L. mitchellae E. A. Smith, 1900, Sarawak (BMNH) (shell height 15 mm). See also figs. 6a-b, 7a-b. 150 BASTERIA, Vol. 63, Mo. 4-6, 1999
1262/10); Batu Putih near road Lahad Datu-Sandakan, near Sg. Kinabatangan (V
1469/1); Bt. Gomantong, 30 km S. of Sandakan (leg. Lee et al, UF 114735/3; V 1598/
2); Bt. Kipangi 2 mile N.E. of Sukau (leg. Lee et al., UF 114766/10). Tawau Zone: N.W. of road Lahad Datu-Sandakan crossing with Sg. Segama (V 1650/1); 'Kirk's Cave', 8 km N. ofLahad Datu (V 1247/1); Batu Tenggar, Segarong Hills 25 km E.S.E. ofKunak
(V 1797/1). "British North Borneo" (BMNH/2, see above). "Sabah" (leg. Kavelaar,
RMNH/>10). KALIMANTAN. Kalimantan Timur: 30 km W. of Balikpapan (leg. Van Balgooij, V 2455/1); surroundings Lempake (leg. WWF-IP Forest Fire Project 1998, V
5978/1). Material from elsewhere. Philippines. Palawan (leg. Ridsdale, V 2317).
Shell white, usually with translucent banding parallel to the growth lines, sometimes with of brown below the a transverse zig-zag pattern pale most conspicuously present just suture.
1 Whorls 5 3/4-6 /2, the first few moderately convex, the others almost flat to slightly convex, but the last also slightly to moderately concave towards the periphery; periphery with a distinct keel that rather abruptly changes into a less conspicuous edge in the last 1/3-1/4 whorl, thus disrupting the curvature ofthe keel when the shell is observed from below. Suture moderately depressed between the first whorls, elsewhere slightly depressed. Spiral sculpture: fine combined with 7 Umbilicus a striation, up to slightly raised, inconspicuous ridgelets. open, narrow, partly covered by the peristome. Peristome white, on the palatal side usually slightly thickened and abrupdy reflected, and often with a slight to distinct lip that gradually decreases in height towards the angular edge; parietal side usually thin, but sometimes with low continuation of the the side. 14-21 width 14- a ridge as a lip on palatal Height mm,
24 mm; height aperture 7-12 mm, width 8-15 mm. Animal the shell, therefore, with white green, shining through shell; looking green (opaque) patches in living specimens. and forest various soil in and Ecology. Primary secondary on types, even kerangas peat swamp forest on wet, acid soils. Also found in oil palm plantations, in coastal sand well in forest 3000 asl. Observed shrubland, on dunes, as as montane at up to m living on vegetation. - Distribution. - Borneo: and found in probably widespread fairly common, not Kalimantan Selatan. Also in The Philippines: a record from Palawan (see above), and probably elsewhere but known under a different name.
Notes. Variable in size and the ratio height/width. In some thick-shelled specimens the peripheral keel on the last whorl is shallowly and obtusely serrate.
3- Leptopoma bourguignati Issel, 1874 (fig. 7)
Leptopoma bourguignati Issel, 1874: 65; syntypes ("Territorio di Sarawak") leg. Doria & Beccari, MCSNG/2
(not seen).
Leptopoma (Trocholeptopoma) mitchellae E. A. Smith, 1900: 42; syntypes ("Sarawak, North Borneo") BMNH
1900.1.15.1-2/2.
Material seen. SARAWAK. Ist Div.: Bako N.P. (V 1436/1). "Sarawak, North Borneo" (BMNH/2, see above).
Shell white, with translucent patches or banding obliquely crossing the growth lines.
Whorls 5 3/4-5 5/8, the first few moderately convex, the others slightly convex, but the last also towards the usually slightly concave periphery; periphery with a moderately distinct keel that into less towards the may gradually change a conspicuous edge ap- Vermeulen: Non-marine molluscs of Borneo (Cyclophorus, Lxplopoma & Craspedotropis) 151
6-8. L. Sabah 18 Figs. Leptopoma spec. 6, undalum (Metcalfe, 1851), (RMNH) (shell height mm).
L. ofL. mitchellae E. A. Sarawak 15 7, bourguignati Issel, 1874, syntype Smith, 1900, (BMNH) (shell height mm).
8, L. signatum Pfeiffer, 1856, syntype, “Borneo” (BMNH) (shell height 10.5 mm). 152 BASTERIA, Vol. 63, No. 4-6, 1999
erture, but without disrupting the curvature of the keel when the shell is observed from below. Suture close the moderately depressed, except to aperture. Spiral sculpture: a fine striation, combined with up to 9 slightly raised, inconspicuous ridgelets. Umbilicus
covered the Peristome white, the side not open, narrow, partly by peristome. on palatal reflected without thin side. thickened and gradually to abruptly and a lip; on the parietal 14-21 width 14-24 7-12 width 8-15 Height mm, mm; height aperture mm, mm.
Ecology. One record from podzolic forest on wet, acid soils on sandstone bedrock, at 50 m asl.
Distribution. Borneo: Sarawak, Ith Div.
Notes. Looks like thin-shelled, not entirely adult L. undatum, and is easily confused with that species. The regularly rounded periphery of the last whorl of L. bourguignati offers the best diagnostic character.
4- Leptopoma signatum Pfeiffer, 1856 (fig. 8)
Leptopoma signatum Pfeiffer, 1856: 338; syntypes ("Borneo") leg. Cuming, BMNH 1998014/3.
Cyclostoma signatum Pfeiffer, 1858: 71.
Material seen. - "Borneo" (leg. Cuming, BMNH/3, see above)
Shell without translucent with somewhat corneous, banding, an oblique, zig-zag pattern of brown colouring. Whorls 5-5 1/8, convex, periphery rounded, with a thread-like, rather distinct ridgelet. Suture depressed. Spiral sculpture: some inconspicuous striation, above the periphery also with 4-5 rather inconspicuous, thread-like ridges, below the periphery with such Umbilicus covered 1-3 ridges. open, narrow, partly by the peristome. Peristome white, on the palatal side slightly thickened, rather gradually to rather abruptly reflected
and then distinctly convex, without a lip, abruptly narrowing towards the angular edge.
Height 10-10.5 mm, width c. 10 mm; height aperture 5.5-6 mm, width 5.5-6.5 mm.
Notes. Differs from L. sericatum in having a strongly reflected, convex peristome,
abruptly narrowing towards the angular edge, and in having oblique brown markings on the shell.
The syntype specimens from the Cuming collection are the only shells available. mistakes in the of the collection makes their Frequent labelling Cuming provenance somewhat uncertain.
5- Leptopoma pellucidum (Grateloup, 1840) (fig. 10)
Cyclostoma vitrea Lesson, 1831: 346; type ("Nouvelle Guinee") (not seen). Not Cyclostoma vitrea Draparnaud,
1801; see Forcart, 1952: 85.
Cyclostoma pellucida Grateloup, 1840: 169; type ("Philippines, Manille") (not seen)
Leptopoma vitreum; Pfeiffer, 1847: 108.
Leptopoma pellucidum; Pfeiffer, 1847: 108.
bicolor unknown. A label with the Cyclostoma Pfeiffer, 1852b: 145; syntypes (locality specimens states"Borneo")
leg. Cuming, BMNH 1998012/3.
Leptopoma bicolor; Pfeiffer, 1852: 104.
1855: of BMNH Leptopoma lowi Pfeiffer, 210; type ("Isle Labuan") leg. Low, (not seen).
Leptopoma bicolor anastomoticum Von Martens, 1908: 278. Type ("Samarinda") (not seen). Vermeulen: Non-marine molluscs of Borneo (Cyclophorus, Leptopoma & Craspedotropis) 153
Material SARAWAK. 4th Div.: G. Subis seen. (Batu Niah) (leg. Dorman et al, UF 196485/2; V 1535/2, see note 3 below). BRUNEI. "Brunei" (leg. v.d. Poll, RMNH/
6). SABAH. Kudat Zone: P. Banggi, southernmost point (V 1446/7). Sandakan Zone:
Bt. Gomantong, 30 km S. of Sandakan (leg. Lee et al, UF 114722/3; V 1597/6); Bt.
Kipangi 2 mile N.E. of Sukau (leg. Lee et al., UF 114757/1). Tawau Zone: N.W. of road Lahad Datu-Sandakan crossing with Sg. Segama (V 1651/>10); 'Kirk's Cave', 8 km N. of Lahad Datu (V 1251/4); Batu Tenggar. Segarong Hills 25 km E.S.E. ofKunak (V 1800/>10); Bt. Pababola, Segarong Hills 25 km E.S.E. of Kunak (V 1751/4).
KALIMANTAN. Kalimantan Selatan: Batu Apoh, approximately 35 km N.E. of Martapura (V 3328/1; leg. Lamb & Mackinnon, V 2461/1); Beramban, approximately
13 km E. of Rantau (V 3756/1); Bt. Pagat, limestone hill 8 km S.E. of Barabai (V 3783/ G. 2); Siamang near Desah Liu, 30 km E. of Tandjung (V 3083/1); G. Buleh, 4 km E. of Muara 3687/10 Muara limestone Uja (V ); Jaro near Uja, escarpment W. of the
7 km N. of Kin limestone W. of km village (V 3188/>10); tap (V 3666/1); escarpment 6 road Benualawas-Batulicin limestone road Benuala- (V 3406/5); escarpments along was-Limbungan (V 2843/>10). Kalimantan Timur: Samarinda (leg. v. Roon, RMNH/ Batu Muara main road 3); Butuk, near Komang along Banjarmasin-Balikpapan (V 3518/>10); G. Melihat, foot of S. facing slope, between river and main road Banjar- masin-Balikpapan (V 2955/>10); road Bontang-Sangatta, 6 km N. ofjunction with road into Kutei National Park (leg. Van Valkenburg & Galzin, V 4518/>10); Kutei National
Park (leg. WWF-IP Forest Fire Project 1998, V 5932/1); Pt. ITCI logging concession Van & V G. Sekarat Van (leg. Valkenburg Galzin, 4527/1); Sangkulirang area, (leg. Valkenburg & Galzin, V 4504/2; do, V 4530/3); surroundings Lempake (leg. WWF- IP Forest Fire V "Borneo" Project 1998, 5976/3). (leg. Cuming, BMNH/3, see above; leg. Schwaner, RMNH/2).
Shell white to corneous, sometimes with a pale brownish-purple apex, with or without translucent banding parallel to the growth lines as well as following the most prominent spiral sculpture; often with a transverse zig-zag patternof brown colouring, or with thin, brown spiral bands following the most prominent spiral ridges, a few of these sometimes fused to wider colour bands; shells with combined transverse and spiral colour patterns also Whorls rounded occur. (4 3/8-)5-5 1/2, convex, periphery (rarely angular, see note
3, below), at most with a thread-like, inconspicuous ridgelet. Suture depressed. Spiral fine above the often with five similar sculpture: a striation, periphery up to ridgelets, to the peripheral ridgelet but weaker, below the periphery sometimes one such a ridge- let. Umbilicus open, narrow, at most partly covered by the peristome. Peristome white, on the palatal side usually slightly thickened, abruptly reflected but then about flat, with without or a slight lip, gradually narrowing towards the angular edge; parietal side thin. 10.2-16 width 10.2-16 shell often than usually Height mm, mm, slightly higher wide; height aperture 5.8-9.2 mm, width 6.2-9.5 mm. Forest limestone soil. No certain records from other bedrock Ecology. on types on
Borneo. Extralimitally observed living on the forest floor as well as on vegetation.
th Distribution. Borneo: widespread but not known from Sarawak, the 4 Division Extralimital due excepted. range unknown to uncertain taxonomy, material assigned to L. and its L. vitreum forms with distinct pellucidum synonym (excluding spiral sculpture, which are included here in L. sericatum) is known from Taiwan southwards to Papua New
Guinea, as well as from Indonesia, Malaysia and Indochina. - Notes. Leptopoma pellucidum constitutes an unresolved species complex; extreme in and have variability occurs size, spiral sculpture colouring. Specific names been proposed for various forms throughout its range. Unequivocal distinction between these 154 BASTERIA, Vol. 63, No. 4-6, 1999
forms, however, appears impossible because intermediates exist, and because the states of the above characters in almost combination. A broad occur every possible species concept has been applied here, although this is unsatisfactory because some forms are restricted small of the of the to a part range species. Bornean L. pellucidum differs from L. sericatum mainly in the spiral sculpture. Where occurring together, the two forms can usually be separated unequivocally, but interme- diate specimens occur occasionally. Because of the distributional pattern the forms cannot be given subspecific rank. Therefore, they are provisionally classified as separate species here.
A population at G. Subis (Batu Niah, the only known from Sarawak) differs from L. pellucidum elsewhere on Borneo in having more slender shells and an angular periphery.
Because they are morphologically enclosed within the species complex, they have not been given taxonomic rank.
6- Leptopoma sericatum (Pfeiffer, 1851) (fig. 9)
Cyclostoma sericatum Pfeiffer, 1851: 244; syntypes ("North Borneo") leg. Taylor, colln. Cuming, BMNH
1998013/5).
Leptopoma sericatum; Pfeiffer, 1852: 108
Godwin 1889: 337. Material colln. Leptopoma wallacei auct. Austen, ("Borneo") Hungerford (BMNH, not seen). Not Cyclostoma (Leptopoma) wallacei Pfeiffer, 1857, from the Aru Islands.
Leptopoma sericatum var. baramense Kobelt, 1897: 29; lectotype ("Baramfluss") leg. Kiikenthal, SMF 126429
(not seen).
Leptopoma (sericatum var.) baramense; Kobelt, 1902b: 409
Leptopoma sericatum baramense; Zilch, 1954: 153.
Material seen. SARAWAK. Ist Div.: near Bau (leg. Lim Chan Koon, V 5803/
4; leg. De Vogel, V 2625/1); 2-3 km S.W. of Bau (leg. Dorman, UF 196277/1); G. Kapur 6 km S.E. of Bau (V 2245/6); Kpg. Tiang Bekap 10 km S.S.W. of Kpg. Beratok
(leg. De Vogel, V 2572/2); G. Selabor, W. of Kpg. Lobang Batu, 12.5 km S. of
Tebakang (V 2092/3). 4th Div.: G. Subis (Batu Niah) (leg. Dorman et al., UF 196415/ 1; do., UF 196453/1; do., UF 196484/1; V 1536; V 1543; leg. Schuiteman, V 5456);
do., W. ofBatu Niah (leg. Raven, R L1628/>10); do., 4 km S. of Batu Niah (leg. Raven,
R L 1662); Lambir Hills (leg. Raven, R/l); G. Pala near Batu Bulang, near G. Mulu
National Park (leg. Schuiteman, V 5400/2); G. Mulu National Park, near National Park entrance (leg. Ball, V 2674/2); do., trail to G. Mulu (leg. Schuiteman, V 4236/3); do.,
Sg. Melinau Paku headwaters, G. Api (leg. Schuiteman, V 5659/2). BRUNEI. Tem- burong, Labi Road 13 km from Sg. Liang (leg. De Vogel, V 2492/3). SABAH. Interior
Zone: Lian Cave 12 km N. of Keningau (V 1130/1); Batu Punggol S.E. of Sepulot (leg.
Dorman, UF 196629/5; V 1899/2); 0.6 km S. of Kpg. Simatuoh, on Simatuoh Creek
(leg. Dorman, UF 196743/1); hill 3 km ENE. of Simatuoh, 0.5 km E. of Sg. Sapulot
(leg. Dorman, UF 196595/1); 1 km S.E. of Simatuoh, 10 km S.E. of Sepulot (leg. Dorman, UF 196567/6); 5.5 km NNE. of Simatuoh, 9 km E. of Sepulot (leg. Dorman,
UF 196517/2); Pun Batu approximately 30 km W. of Sepulot (V 1297/5). West Coast
Zone: P. Tiga (RMNH/2); P. Gaya (leg. Kittel, K/l); G. Kinabalu National Park, Silau-Silau Headquarters area (leg. Kittel, V 4802/1); do., trail (leg. Kittel, K/4); do., Liwagu (leg. Kittel, K/6); do., Carson Falls (leg. Kittel, K/l). Sandakan Zone: Bt.
Gomantong, 30 km S. of Sandakan (leg. Lee et al., UF 114713/1; do., UF 114722/ Vermeulen: Non-marine molluscs of Borneo (Cyclophorus, & Craspedotropis) 155
12 Figs. 9-10. Leptopoma spec. 9, L. sericatum (Pfeiffer, 1851); a-b, Sarawak, G. Subis (V) (shell height mm); c,
Sabah, G. Madai 40 km SSW. of Lahad Datu (V) (shell height 13 mm). 10, L. pellucidum (Grateloup, 1840); a-b, Sabah,NW. ofroad Lahad Datu-Sandakan crossing with S. Segama (V) (shell height 13 mm); c-d, Sarawak,
G. Subis (V) (shell height 16 mm); d, operculum. 156 BASTERIA, Vol. 63, No. 4-6, 1999
1; V 1595/1); Sukau, Sg. Mananggol (leg. Chan, V 4238/1; Bt. Kipangi 2 mile N.E.
of Sukau (leg. Lee et al, UF 114757/>10); Kinabantangan Valley (leg. Chan, V 5596/ 1). Tawau Zone: Kirk's Cave', 8 km N. of Lahad Datu (V 2520/2); G. Madai, 40 km
S.S.W. of Lahad Datu (V 1710/8); G. Baturong, 50 km W.S.W. of Lahad Datu (leg. Dorman, UF 196778/2; do., UF 196791/4; V 1839/6). "North Borneo" (leg. Taylor, colln. colln. Cuming, BMNH/5, see above; Venmans 5401, RMNH/2). KALIMAN- TAN. Kalimantan Barat: Dutch Borneo 1893-94, Liang Kubung (leg. exp. RMNH/4);
Dadap Ketungau (do., RMNH/2)! Kalimantan Selatan: Nateh near Batu Tangga,
approximately 18 km E. of Barabai (V 3019/1); G. Siamang near Desah Liu, 30 km
E. of near Muara limestone W. of the Tandjung (V 3084/>10); Jaro Uja, escarpment limestone road village (V 3183/2); escarpments along Benualawas-Limbungan (V 2842/ 1). Kalimantan Timur: Kutei National Park (leg. WWF-IP Forest Fire Project 1998, V 5933/1); surroundings Merabu: Liang Belana (do., V 5942/1); surroundings Lempake
(do., V 5977/3).
As the nominate subspecies, but also with a pinkish apex; without transverse zig-zag colour often with brown thin pattern, orange or pinkish spiral bands following the most
prominent spiral ridges, the lowermost of these sometimes fused to wider colour bands. Whorls 3/8-5 4 l/8(-5 1/2), periphery rounded to angular, usually with a moderately
distinct thread-like ridge. Spiral sculpture: a fine striation, above the periphery with 3-
6 rather distinct, thread-like ridges, below the periphery l-2(-6) such ridges. Height 9-15 width 9.2-15.5 shell often wider than mm, mm, slightly high; height aperture 5-8 mm,
width 5-9.5 mm.
Animal red, tips of tentacles blackish (1 observation).
Ecology. In primary forest on limestone soils, also in forest on soils over sandstone/
shale bedrock, and granodiorite. Also found in disturbed vegetation, and in roadsides.
montane at Found in lowland conditions, but also in forest about 1800 m asl. Observed
living on the forest floor as well as on vegetation.
Distribution. Borneo: Extralimital L. altum Moellen- widespread. range unknown;
dorff, 1897, from Java, and L. fultoni Aldrich, 1898, from Sumatra, are possibly identical,
as well as some species described from the Philippines.
Species excluded from the list of Bornean Leptopoma.
Leptopoma duplicatum auct.
L. Leptopoma duplicatum auct. Bock, 1881: 634; material ("Mindai")(not seen). Possibly not duplicatum Pfeiffer,
1856, of unknown origin.
Leptopoma massena auct.
Leptopoma massena auct. Bock, 1881: 634; material ("Mindai") (not seen). Probably not Cyclostoma massena
Lesson, 1831, from New Guinea.
Leptopoma subconicum auct.
Leptopoma subconicum auct. Bock, 1881: 634; material ("Mindai") (not seen). Probably notCyclostoma subconica
Pfeiffer, 1862, probably from southern Vietnam. Vermeulen: Non-marine molluscs of Borneo (Cyclophorus, Leptopoma & Craspedotropis) 157
Notes. The identifications of Bornean Leptopoma in Bock (1881: 634) are probably not either of correct: the species concerned are not likely to occur on Borneo, or they are doubtful and taxonomic status. The shells available for and provenance are not study, cannot be identified. At most, their identity can be inferred from checking properly identified material of the species listed above.
Both L. duplicatum auct., and L. subconicum auct., probably refer to shells combining thread-like with brown colour could be L. but also spiral ridges a pattern. They signatum, various species of the genus Japonia, in which this combinationof characters is common.
Leptopoma massena auct. probably refers to a shell of L. pellucidum.
Leptopoma tenebricosum (Adams & Reeve, 1850)
Cyclostoma tenebricosum Adams & Reeve, 1850): 57. Type ("Balangbanganinsulae Borneo") leg. Cuming (not seen)
Leptopoma tenebricosum Pfeifler, 1852: 117.
Cyclophorus tenebricosus Pfeifler, 1865: 96.
uncertain. Could be either Note. Identity a Leptopoma, or a Japonia.
Leptopoma whiteheadi E. A. Smith, 1887.
BMNH 89.5.21.2- Leptopoma whiteheadi E. A. Smith, 1887: 133; syntypes ("Northern Borneo") leg. Whitehead,
3/2.
Material seen. "Northern Borneo" (BMNH/2, see above).
Note. - This is a Japonia species
Craspedotropis Blanford, 1864
Shell small somewhat with very (1.8-3.6 mm wide), thin, translucent, a silky lustre, whitish without colour Protoconch transition to corneous, markings. finely rugose, to-
wards the teleoconch unmarked. Spire conical to high conical. Radial sculpture: growth
lines and fine riblets. Spiral sculpture: striation, often combined with ridges. Umbilicus
covered the often with one or more Peristome open, not by peristome. Aperture edges. without sinus. Periostra- somewhat thickened but not reflected, simple; angular edge a
cum thin, corneous, with or without hairs or other appendages. with inner the of the whorls Operculum thick, many whorls, layer corneous, margins
erect, the spiral cavity in between filled with calcareous matter or not. Distribution. India, Borneo. - - Notes. Three genera seem appropriate to include the species listed below: Cya- thopoma, Blanford, 1861 (India to Africa), Jerdonia Blanford, 1861 (India mainly), and
Craspedotropis Blanford, 1864 (India). According to Kobelt (1902), the first two have an
operculum with calcareous elements, whereas the third has an entirely corneous oper-
erect culum.Cyathopoma should differfromJerdonia in having an operculum with free and
margins. Gude (1921) is less explicit about the generic delimitations, but, judging from the species descriptions, follows Kobelt. 158 BASTERIA, Vol. 63, No. 4-6, 1999
of material in However, on investigation present RMNH, Craspedotropis cuspidata (Ben-
the of the to have a son, 1851), type species genus, appears corneous, multispiral operculum with thin, free and erect whorl margins. The spiral cavity in between the filled of This is similar margins is partly with a layer calcareous matter. fairly to Cya-
and two thopoma, the only remaining distinction between the genera then lies in the shape of the shell: Craspedotropis usually has a higher conical spire and has a distinct peripheral keel, or a number of equally distinct ridges. All in all, the distinctness of the three genera inclusion of the below in of them is leaves much to be desired, and the species any doubtful. The argumentation to choose Craspedotropis is as follows: C. andrei, is included because the shell is similar that of C. the of the Unfor- shape to cuspidata, type genus. C. tunately, the operculum of andrei is unknown.Craspedotropis borneensis has an operculum essentially similar to that of C. cuspidata, although it lacks the calcareous layer. The shell shape is reminiscent of Indian species later included in the genus: C. bilirata (Beddome, 1875), and C. salemensis (Beddome, 1875). Craspedotropis juvenilis, finally, is in fact fairly similar to C. andrei, except for the prominent peripheral keel of the latter. However, it would equally well fit in Cyathopoma as far as the shell is concerned. Again, the operculum
is lacking in the only specimen available. Next the listed the includes four from India 1921: to taxa below, genus species (Gude,
15), and probably one more from Bunguran Isl. off the NW. point of Borneo: Cyathopoma tricarinatum E. A. Smith, 1894.
Bornean Japonia, Gould, 1859 (is Lagocheilus, Blanford, 1864) have larger shells (the
smallest about 4 mm high), usually have a reflected peristome with a sinus in the angular edge, and have an operculum without erect whorl margins. It is either entirely corneous, thin of side. or it has a layer calcareous matter on one
Key to the Bornean species ofCraspedotropis
(check as large series of specimens as possible)
la Ultimate whorl entirely without conspicuous ridges 1- Craspedotropis juvenilis
1 b Ultimate whorl with a distinct peripheral keel, or with various distinct spiral ridges 2
2a Ultimate whorl with 9 about equally distinct spiral ridges 2- Craspedotropis borneensis
with 2b Ultimate whorl a single peripheral keel, all other spiral sculpture inconspicuous
3- Craspedotropis andrei
1- Craspedotropis juvenilis spec. nov. (fig. 11)
Material seen. SARAWAK. 4th Div.: G. Mulu National Park, Sg. Melinau Paku
headwaters, G. Api (leg. Schuiteman, V 5430/1, holotype RMNH 59386).
Spire high conical. Whorls 3 5/8, convex; periphery rounded. Suture depressed. Radial sculpture: inconspicuous growth lines, alternating with somewhat spaced riblets
(c. 20/mm on the last whorl). Spiral sculpture: inconspicuous striation, alternating with fine 12 the last shell with fine reticulate Umbilicus ridgelets (c. on whorl); a very sculpture.
narrow. Aperture slightly angular above, otherwise circular. Peristome with a thickened the side. 1.9 width 1.8 and width 0.95 lip on palatal Height mm, mm; height aperture
mm. Periostracum greenish, smooth, without hairs or appendages.
Ecology. Forest on limestone soil. Found at 1200 m asl. Distribution. Borneo: G. Sarawak, 4th Div., Mulu area only Vermeulen: Non-marine molluscs of Borneo (Cyclophorus, Leptopoma & Craspedotropis) 159
Notes. - - The low whorl this immature The count gives species an appearance. is peristome, however, thickened, as in adult shells of most cyclophorids. Its inclusion in is somewhat from the absence of Craspedotropis doubtful, but, apart a
peripheral keel, the shell is more similar to C. andrei than to any other Bornean cyclo- phorid.
2- Craspedotropis borneensis (Godwin Austen, 1889) (fig. 12)
Jerdonia? borneensis Godwin Austen, 1889: 345; holotype ("Busan Hills") leg. Everett, BMNH 1889.12.7.21/1
Material SARAWAK. seen. Ist Div.: Busau (leg. Everett, BMNH/1, see above).
Spire high conical. Whorls 5 1/2, convex. Suture depressed. Radial sculpture:
growth lines and densely placed riblets (35/mm on the last whorl) which are most
to suture. distinct close the Spiral sculpture: distinct, high ridges, 9 on the last whorl
(6 regularly spaced on the lateral surface of the whorl, 2 on the basal surface, but at
some distance from the lowermost lateral one, and 1 in the umbilicus); in between
the ridges a fine striation that is least conspicuous near the suture. Umbilicus rather
wide. less with shallow sinus Peristome with Aperture more or ovate, a basally. a slight the and the basal distance from lip on palatal side, at some the margin. Height 3.2 width 1.6 1.2 width 1.3 Periostracum mm, mm; height aperture mm, mm. greenish, close the raised to suture to minute crests over the radial riblets, without further appendages. free widened Operculum corneous, whorl-margins and flattened so that together they
form a second corneous layer at some distance from the first. No calcareous matter
present.
Ecology. Forest on limestone soil. Lowland conditions.
Distribution. Borneo: Sarawak, Ist Div., limestone ranges S.W. of Kuching.
3- andrei Craspedotropis spec. nov. (fig. 13)
Material seen. SARAWAK. 4th Div.: Bt. Pelamau (leg. Haegens, V 5614/1); G.
Mulu National Park, Sg. Melinau Paku headwaters, G. Api (leg. Schuiteman, V 5431/ 2, including holotype RMNH 59385; do., V 5640/3).
conical. Whorls the first few Spire 4-4 1 /4, convex, the last almost flat to moderately
convex, slightly concave towards the periphery; periphery with a distinct keel. Suture
depressed. Radial sculpture: growth lines and densely placed to irregularly spaced riblets
(15-25/ mm on the last whorl) which are most distinct close to the peripheral keel. Spiral
sculpture: inconspicuous striation, combinedabove the periphery with 4-5 slightly raised,
somewhat more distinct ridges; below the periphery with 1-2 such ridges, one of these
running close to the peripheral keel. Umbilicus rather wide. Aperture angular above and the Peristome to right. with a thickened lip on the palatal and basal side. Height 2.0- 2.5 width 2.8-3.6 1.0-1.1 width 1.3-1.6 Periostracum mm, mm; height aperture mm, mm.
corneous, sometimes raised to a minute crest where radial riblets cross the spiral ridges,
with rather deciduous, distinct, flat, obtuse appendages of c. 0.1 mm long on the
peripheral keel. 160 BASTERIA, Vol. 63, No. 4-6, 1999
Figs. 11-12. Craspedotropis spec. 11, C. juvenilis spec. nov., holotype; Sarawak, G. Mulu area, G. Api (RMNH)
(shell height 1,9 mm). 12, C. borneensis (Godwin Austen, 1889), holotype; Sarawak, Busau (BMNH); a-b, shell
3.2 lateral section whorls. (height mm); c-e, operculum; c, outside; d, view; e, through outer Vermeulen: Non-marine molluscs of Borneo (Cyclophorus, Leptopoma & Craspedotropis) 161
13. andrei G. Mulu Fig. Craspedotropis spec. nov. a-b, holotype, Sarawak, area, G. Api (RMNH) (shell height
2.2 Bt. Pelamau mm); c-d, paratype, Sarawak, (V) (shell height 2.4 mm). 162 BASTERIA, Vol. 63, No. 4-6, 1999
Ecology. Forest on limestone soil. Found at 1200-1300 m asl.
Distribution. Borneo: Sarawak, 4th Div., G. Mulu area and surrounding limestone hills.
Note. Named in honour of Mr. Andre Schuiteman (Rijksherbarium, Leiden, the Netherlands), who collected the first specimens of this species, and whose collecting activities at 900-1200 m asl. in the Pinnacle area in the Mulu range revealed the extraordinary composition of the local snail fauna.
ACKNOWLEDGEMENTS
Thanks are due to the Nationaal Natuurhistorisch Museum, for the use of the library and other facilities Dr. A. C. has read the put at my disposal. van Bruggen carefully manuscript. The Florida Museum of Natural History, Gainesville, sent a large amount material available of unidentifiedmaterial on loan. More was kindly made by the British
Museum (Natural History), London. Interesting material was also obtained on loan from the private collections of Dr. J. G. M. Raven and Mr. K. Kittel.
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